Facultative amphidromy involving estuaries in an annual amphidromous fish from a subtropical marginal range

Variation in the life cycle of diadromous fishes can be explained by differential food availability between marine and freshwater habitats, since migration is often interpreted as a mechanism for exploiting food resources. Theoretically, a migration pattern of mainly remaining in freshwater occurs in tropical and subtropical habitats where fluvial productivity possibly exceeds marine productivity. However, in Yakugachi River, Amami-Oshima Island, southern Japan, low nutrient concentrations in the river suggest that food availability is limited for the subtropical Ryukyu-ayu Plecoglossus altivelis ryukyuensis. Since Ryukyu-ayu is an amphidromous fish that mainly grows in rivers after spending 2 months in the sea, limited food availability in rivers would force this species to migrate to other habitats with better food availability. Otolith increment and Sr:Ca analyses of 48 adult Ryukyu-ayu collected from the Yakugachi River revealed that all individuals visited estuaries more than three times after moving upstream. Although the specific growth rates of this species in the river had no correlation with the salinity profile in the fluvial period, this movement may be an adaptive choice because the salinity profile significantly affected the body size at maturity. Our results highlighted individual-based variations in amphidromous migration for utilizing estuaries, which could be explained by relatively higher productivity in estuarine than in freshwater and marine habitats.     

Reconstructing larval growth and habitat use in an amphidromous goby using otolith increments and microchemistry

High‐resolution analysis of growth increments, trace element chemistry and oxygen isotope ratios (δ¹⁸O) in otoliths were combined to assess larval and post‐larval habitat use and growth of Awaous stamineus, an amphidromous goby native to Hawai‘i. Otolith increment widths indicate that all individuals experience a brief period of rapid growth during early life as larvae and that the duration of this growth anomaly is negatively correlated with larval duration. A protracted high‐growth period early in larval life is associated with a lower ratio of Sr:Ca, which may reflect low salinity conditions in nearshore habitats. A distinct shift in δ¹⁸O (range: 4–5‰) is closely associated with the metamorphic mark in otoliths, indicating that larval metamorphosis occurs promptly upon return to fresh water. Strontium and other trace elements are not as tightly coupled to the metamorphosis mark, but confirm the marine‐to‐freshwater transition. Integration of microstructural and microchemical approaches reveals that larvae vary substantially in growth rate, possibly in association with habitat differences. Although time and financial costs make it difficult to achieve large sample sizes, present results show that examining even a small number of individuals can lead to novel inferences about early life history in diadromous fishes and illustrates the value of integrating analyses.     

Early starvation limits survival in amphidromous fishes

Newly hatched amphidromous larvae drift downstream to the sea. Since there is practically no food available for drifting larvae, an extended downstream migration may enhance the risk of starvation. We hypothesized that early survival of amphidromous fish varies among populations according to the length of the river, due to its effect on larval starvation. The hypothesis was tested with the common Japanese amphidromous goby, Rhinogobius brunneus . Based on morphological characteristics, larvae were categorized as being in an endogenous feeding state (EFS) or irrecoverable starved state (ISS). The frequency of ISS larvae in the population from a long river was nearly twice the frequency in the population from a short stream. More than half of larvae were judged to be dying during downstream migration in the comparatively long river, although this was possibly an underestimation. It is suggested that early starvation limits reproductive success far from the sea.     

Validation and efficacy of transgenerational mass marking of otoliths in viviparous fish larvae

Transgenerational mass marking of viviparous fish larvae in vivo was validated by intra‐muscular injection of elemental strontium chloride (SrCl₂) in gestating females and detection of the Sr in the otoliths of developing larvae. All otoliths of brown rockfish Sebastes auriculatus larvae produced from SrCl₂‐injected females showed enriched Sr:Ca ratios near the otolith edges, and the signatures did not appear to be affected by the anterior, centre and posterior positions of larvae within the ovary. Results from the present study indicate that transgenerational marking is a highly reliable technique for marking large numbers of extremely small viviparous fish larvae.     

Migration flexibility between freshwater and marine habitats of the pond smelt Hypomesus nipponensis

The life histories of the pond smelt Hypomesus nipponensis collected from Japanese fresh waters and brackish (sea) waters were studied by examining the strontium (Sr) and calcium (Ca) concentrations in their otoliths. The Sr:Ca ratios in the otoliths changed with the salinity of the habitat. The pond smelt living in a freshwater environment showed consistently low Sr:Ca ratios throughout the otolith, averaging 1·2–1·3 × 10⁻³. These samples were identified as a standard freshwater type. In contrast, fish collected from the intertidal zone showed higher otolith Sr:Ca ratios than those in the standard freshwater type, and the ratios fluctuated along the growth phase. In addition to the two representative life‐history types of H. nipponensis , i. e . freshwater and anadromous life‐history types, other pond smelts were found to have an estuarine resident life history‐type with no freshwater phase, indicating that the pond smelt has a flexible migration strategy with a high degree of behavioural plasticity and an ability to utilize the full range of salinity in its life history.     

Flexible Life History Strategies of Ninespine Sticklebacks,Genus <Emphasis Type="BoldItalic">Pungitius</Emphasis>

Synopsis We studied the life histories of the ninespine sticklebacks, Pungitius pungitius and Pungitius tymensis, collected from Japanese freshwater and brackish (sea) water habitats by examining the strontium (Sr) and calcium (Ca) concentrations in their otoliths. The Sr:Ca ratios in the otoliths changed with the salinity of the habitat regardless of identification as freshwater or brackish water type based on morphological characteristics. The ninespine sticklebacks living in a freshwater environment showed consistently low Sr:Ca ratios throughout the otolith. These samples were identified as a standard freshwater type. In contrast, all freshwater-type fishes collected from the intertidal zone showed higher otolith Sr:Ca ratios than those in the standard freshwater type, and the ratios fluctuated with the growth phase. All brackish water-type fishes collected in the intertidal zone showed the highest otolith Sr:Ca ratio throughout the otolith. In the present study, besides the two representative life history types of P. pungitius, i.e., freshwater and brackish water life history types, other sticklebacks had an anadromous life history type. These findings clearly indicate that the ninespine stickleback has a flexible migration strategy with a high degree of behavioral plasticity and an ability to utilize the full range of salinity in its life history.     

Otolith microchemistry of tropical diadromous fishes: spatial and migratory dynamics

Otolith microchemistry was applied to quantify migratory variation and the proportion of native Caribbean stream fishes that undergo full or partial marine migration. Strontium and barium water chemistry in four Puerto Rico, U.S.A., rivers was clearly related to a salinity gradient; however, variation in water barium, and thus fish otoliths, was also dependent on river basin. Strontium was the most accurate index of longitudinal migration in tropical diadromous fish otoliths. Among the four species examined, bigmouth sleeper Gobiomorus dormitor, mountain mullet Agonostomus monticola, sirajo goby Sicydium spp. and river goby Awaous banana, most individuals were fully amphidromous, but 9–12% were semi‐amphidromous as recruits, having never experienced marine or estuarine conditions in early life stages and showing no evidence of marine elemental signatures in their otolith core. Populations of one species, G. dormitor, may have contained a small contingent of semi‐amphidromous adults, migratory individuals that periodically occupied marine or estuarine habitats (4%); however, adult migratory elemental signatures may have been confounded with those related to diet and physiology. These findings indicate the plasticity of migratory strategies of tropical diadromous fishes, which may be more variable than simple categorization might suggest.     

Unravelling the life-history patterns and habitat preferences of the Japanese eel (Anguilla japonica) in the Pearl River,China

Eels have fascinated biologists for centuries due to their amazing long-distance migrations between freshwater habitats and very distant ocean spawning areas. The migratory life histories of the Japanese eel, Anguilla japonica, in the waters of south China are not very clear despite its ecological importance, and the need for fishery regulation and management. In this study, strontium (Sr) and calcium (Ca) microchemical profiles of the otoliths of silver eels were measured by X-ray electron probe microanalysis based on data collected from different habitats (including freshwater and brackish habitats), in the large subtropical Pearl River. The corresponding habitat preference characteristics were further analysed using redundancy analysis (RDA). A total of 195 Japanese eels were collected over 6 years. The collected individuals ranged from 180 to 771 mm in total length and from 8 to 612 g in body weight. Two-dimensional pictures of the Sr:Ca concentrations in otoliths revealed that the A. japonica in the Pearl River are almost entirely river eels, spending the majority of their lives in fresh water without exposure to salt water, while the catadromous migration time has delayed about 1 month in the Pearl River estuary in the past 20 years. RDA analysis further indicated that juveniles and adults preferred water with high salinity and high tide levels. Youth preferred habitats with high river fractals. Our findings contribute to a growing body of evidence showing that the eels are extremely scarce currently and conservation measures against them are imminent, including the protection of brackish and freshwater areas where they live in south China.     

Anadromy and heterogenous population of a tropical shad Tenualosa ilisha in Malaysia,as revealed by otolith microchemistry and molecular evidence

Tenualosa ilisha was found recently in the Perak River in western Peninsular Malaysia. Molecular phylogenetic and haplotype network analyses suggest that T. ilisha has two genetically distinct populations/groups: (i) Peninsular Malaysia (Malaysia population), and (ii) Peninsular Malaysia, Thailand, India and Bangladesh (Indian Ocean population). The results also suggest that the T ilisha population in Peninsular Malaysia is genetically heterogeneous with a typical anadromous migration pattern.     

Retrospective approach to investigating the early life history of a diadromous fish: allis shad Alosa alosa (L.) in the Gironde–Garonne–Dordogne watershed

In this study, the identification of major ontogenetic events and the duration of the fresh water and estuarine phases before the seaward exit of young allis shad Alosa alosa were investigated retrospectively, using the otoliths of spawners. Daily growth increment periodicity in the otoliths was validated with an accuracy of 4 days (9·4%) for fish >30 days. The specific limits of the Sr:Ca ratios in the otoliths, indicating fish movements between salinity 'compartments', were up to 1·1 ×10⁻³ for the freshwater phase, 1·1 to 1·9 × 10⁻³ for the estuarine phase and 1·9 × 10⁻³ for the marine phase. In early life history, a major event occured at 22 days, which was marked on the otoliths by a change in the direction of the otolith growth axis. This event was related to a change in larval swimming behaviour. Juvenile freshwater residence time was between 58 and 123 days, with a median duration of 88 days. Contrary to what has been estimated in previous studies, the present study showed that the estuarine phase lasted from 4 to 36 days, with a median value of 11 days. Seaward exit occurred in fish aged from 63 to 150 days, with a median age of 99 days. These results have important implications for the sustainable management of this species.     

Flexible modes of anadromy in Baltic sea trout: making the most of marginal spawning streams

From examination of the ratios of strontium to calcium laid down as a lifetime record in the otoliths of sea trout Salmo trutta from Gotland, Baltic Sea, it was found that: (1) the shortest stream was used mostly by precociously emigrant or coastally hatched spawners; (2) longer streams had more fish that underwent normal smoltification; (3) sea-caught fish were predominantly coastally hatched (presumably near stream mouths). Furthermore, some otoliths showed no evidence of a freshwater history at all, raising the possibility of a contingent of the coastal population that does not depend on riverine spawning. The results emphasize the importance of the coastal zone as natal and early life habitat for sea trout in the Baltic, particularly with respect to a potential change to a warmer climate which may exacerbate conditions within small, ephemeral trout streams.     

Factors influencing Sr/Ca ratios in otoliths of Girella elevata: an experimental investigation

The utility of Sr/Ca ratios in otoliths as indicators of thermal history in fish was investigated for juvenile Girella elevata . There was no direct relationship between ratios of Sr/Ca and temperature of the water, as has been assumed in many previous studies. Sr/Ca ratios did not decrease when water temperature was elevated from 19 to 28° C. Elevation of ambient Sr levels in the sea water caused a significant increase in Sr/Ca ratios in otoliths, and there was a detectable increase in Sr/Ca ratios in otoliths of some individuals fed an Sr-enriched diet. Multiple factors may influence natural Sr/Ca ratios in otoliths of juvenile G. elevata and their interactions make it difficult to interpret the chronology of conditions experienced by an individual fish during early life in the pelagic or benthic environment.     

Patterns of increment width and strontium: calcium ratios in otoliths of juvenile rock blackfish, Girella elevata (M.)

Otoliths of juvenile Girella elevata (M.) were examined to obtain information about the environmental conditions experienced during early life. Patterns of increment deposition and elemental ratios in otoliths were compared in wild fish. A tetracycline experiment indicated that increments were deposited daily in juveniles. Although different patterns in the spacing of increments were found among juveniles collected at different locations and times, the widest increments were always found in the first 40 increments. Strontium: calcium (Sr: Ca) ratios increased with age in the otoliths of most wild G. elevata .
The patterns of increment width and Sr: Ca ratios were not related and, therefore, were probably not under the same relative control by environmental or physiological factors. Although the number of increments can be used to age juvenile G. elevata , the utility of increment widths and Sr: Ca ratios as environmental predictors in this species is questionable without experimental validation.     

Intra-population variation in anadromy and reproductive life span in rainbow trout introduced in the Santa Cruz River, Argentina

Scale patterns, maturational status and otolith microchemistry (strontium to calcium ratios) were analysed in sympatric anadromous and non-anadromous rainbow trout Oncorhynchus mykiss in the Santa Cruz River (Patagonia, Argentina) to investigate the life-history differences of anadromous and non-anadromous lifestyles and the association between maternal origin and progeny life history. The analyses revealed that both forms can give rise to one another, indicating a single population with alternative phenotypes. Anadromous fish smolted at ages 2 and 3 years, matured after 1 to 2 years in the ocean, and survived up to 11 years, spawning up to eight times. Non-anadromous fish survived up to 6 years, spawning up to three times. The extended reproductive life span associated with anadromy in this river suggests that increased energetic and physiological demands associated with ocean migration may not necessarily result in reduced postspawning survival, as has been suggested for salmonids in general. Alternatively, reduction in parity may be regarded as the evolutionary outcome of reproductive traits resulting from the adoption of anadromy ( i.e. augmented reproductive investment) coupled with long-range migrations to and from the ocean. The life-history patterns of Santa Cruz River rainbow trout provide a natural experiment for investigating the evolutionary transition and maintenance of anadromy and non-anadromy within salmonid populations.     

Migratory benthic fishes may induce regime shifts in a tropical floodplain pond

1. Alternative states are a widely recorded phenomenon in shallow lakes, which may shift between turbid‐ and clear‐water conditions. Here, we investigate whether such shifts in a tropical floodplain pond may be related to the effect of the flood pulse regime on the community structures of fish and macrophytes. 2. Using a long‐term data set, we demonstrate how benthic fish migration together with colonisation by submerged plants affected the transition from a turbid to a macrophyte‐dominated state in a floodplain pond without top‐down control. 3. In our study, the turbid state occurred mostly during low water phases and was largely characterised by high values for the biomass of benthic fish, chlorophyll‐a and total phosphorous. 4. During the period of rising water levels, the migration of benthic fish out of the pond occurs simultaneously with the establishment of submerged plants, while water turbidity decreases along with phytoplankton and nutrient concentrations, inducing a clear‐water phase. However, when submerged plants are absent and fish migration is low, a transient state is generated. 5. We suggest that, in contrast to temperate ponds and shallow lakes, where the main driving mechanisms establishing alternative states are related to cascading effects via the food chain, in tropical ponds and shallow lakes it is resuspension of sediments by benthic fish that plays the most significant role in establishing alternative states. However, the effect of the flood pulse regime plays an important role in the temporal dynamics of fish community structure by controlling benthic fish migration.     

Small anadromous Salvelinus malma at the southern limits of its distribution

Anadromy in the Dolly Varden Salvelinus malma at the southern limits of its distribution were examined by an intensive field survey and measurements of otolith Sr and Ca concentrations. Body sizes of anadromous individuals were small, only slightly larger than those of mature parr, and the degree of anadromy was extremely low.     

Diverse migratory histories in a brackish water type of the ninespine stickleback, <Emphasis Type="Italic">Pungitius pungitius</Emphasis>

The life history in a brackish water type of the ninespine stickleback, Pungitius pungitius, was studied by examining the strontium (Sr) and calcium (Ca) concentrations in the otoliths. The fluctuating patterns of Sr/Ca ratios along the life history transect in the otoliths varied widely among fish in spite of their identification as brackish water type as estimated by morphological characteristics. More than 70% fish showed the intermediate otolith Sr/Ca ratio throughout, averaging 5.23–7.71 × 10⁻³. Besides this brackish water resident life history type of P. pungitius, other sticklebacks had anadromous (25%) and freshwater amphidromous (2.5%) life history types. These findings clearly indicate that the migration of the ninespine stickleback between fresh and sea waters is obligatory but facultative having an ability to utilize the full range of salinity in its life history.     

Migratory diversity predicts population declines in birds

Declines in migratory species are a pressing concern worldwide, but the mechanisms underpinning these declines are not fully understood. We hypothesised that species with greater within‐population variability in migratory movements and destinations, here termed ‘migratory diversity’, might be more resilient to environmental change. To test this, we related map‐based metrics of migratory diversity to recent population trends for 340 European breeding birds. Species that occupy larger non‐breeding ranges relative to breeding, a characteristic we term ‘migratory dispersion’, were less likely to be declining than those with more restricted non‐breeding ranges. Species with partial migration strategies (i.e. overlapping breeding and non‐breeding ranges) were also less likely to be declining than full migrants or full residents, an effect that was independent of migration distance. Recent rates of advancement in Europe‐wide spring arrival date were greater for partial migrants than full migrants, suggesting that migratory diversity may also help facilitate species responses to climate change.     

Diverse migratory histories of Japanese Trachidermus and Cottus species (Cottidae) as inferred from otolith microchemistry

The migratory histories of Japanese freshwater sculpins, one Trachidermus and four Cottus species, were studied by examining strontium (Sr) and calcium (Ca) concentrations in their otoliths using wavelength dispersive X‐ray spectrometry on an electron microprobe. The Sr : Ca ratios in the otoliths changed with salinity of the habitat. The otoliths of Cottus nozawae showed consistently low Sr : Ca ratios, with an average of 3·37 × 10^?3 from the core to the edge, suggesting a freshwater resident life cycle. In contrast, the otolith Sr : Ca ratios for Trachidermus fasciatus and Cottus kazika changed along the life history transects possibly in accordance with their migration patterns from sea to fresh water. The ratios of T. fasciatus and C. kazika averaged 5·4 × 10^?3 and 5·3 × 10^?3 respectively, in the otolith region from the core to the points 450–890 μm, and changed to the lower levels, averaging 2·0 × 10^?3 and 2·7 × 10^?3, in the outer otolith region. These data suggest that both the species have a catadromous life cycle. The otoliths of Cottus hangiongensis had low Sr : Ca ratios in the two regions from the core to the points 15–30 μm and the points 415–582 μm to the edge, averaging 2·0 × 10^?3 and 1·9 × 10^?3, with significantly higher ratios in the narrow area between these regions, averaging 4·6 × 10^?3. Similar ontogenetic changes in otolith Sr : Ca ratios were found in the otoliths of Cottus amblystomopsis, suggesting their amphidromous life cycle. These findings suggest that otolith Sr : Ca ratios reflect individual life histories and that Japanese Trachidermus and Cottus species have diverse migratory histories.